Many of the bolded characters in the characterization above are apomorphies of subsets of streptophytes along the lineage leading to the embryophytes, not apomorphies of crown-group embryophytes per se.

All groups below are crown groups, nearly all are extant. Characters mentioned are those of the immediate common ancestor of the group, [] contains explanatory material, () features common in clade, exact status unclear.

Taxa with relatively small flowers that are aggregated into conspicuous flower-like inflorescences (but c.f. Campanulaceae, etc.; see Leins & Erbar 2010: fig 157) are common in all three large orders in this clade, the apices of the petals tend to be pointed (not in Aquifoliaceae), and valvate corollas are common. The feature "small flowers" itself may be assignable to the euasterid node, since the first nodes of both the lamiid and campanulid clades may have this feature.

Changes in seed size may be pegged to this node. Seeds of this clade are generally rather small, perhaps connected with the herbaceous-shrubby habit so common here, and contrast with the larger seeds of most Aquifoliales, Icacinales, etc. (see also Moles et al. 2005a). Endress (2011a) suggested that the inferior ovary so common here (but see below) might be a key innovation.

The position of early initiation of the corolla tube on the tree is uncertain. Although a number of families show this kind of initiation, not only is sampling within the larger orders poor and within the smaller orders largely non-existent, in the lamiids both Oleaceae and Rubiaceae, basal or almost so in their orders (Lamiales and Gentianales respectively), may have early initiation, and corolla initiation in Aquifoliales and basal clades in the lamiids is largely unknown (Leins & Erbar 2003b for a summary). Furthermore, there may be a developmental connection between early corolla tube formation and the way inferior ovaries in the campanulids develop (Ronse Decraene & Smets 2000). Taxa with more or less superior ovaries are scattered throughout the campanulids, e.g. Sphenostemon (Paracryphiaceae, Paracryphiales), Pittosporaceae (Apiales), Rousseaceae-Carpodetoideae and Phellinaceae (Asterales), within Bruniales, etc., while in Adoxaceae, sister to other Dipsacales, the ovary is semi-inferior, Ying et al. (1993) even describing the ovary of Tetradoxa as being superior, so it will be interesting to see if corolla development changes accordingly.

Chemistry, Morphology, etc. Acetylenic fatty acids/polyacetylenes are sporadic in this clade (e.g. Erbar & Leins 2004; Leins & Erbar 2004b), occurring in Asterales, Dipsacales and Apiales, but always in much embedded clades. These similarities are probably parallelisms. Although iridoids and polyacetylenes usually do not co-occur, the two may be found together, as in Torricellia angulata (Pan et al. 2006; Liang et al. 2009).

The I copy of the RPB2 gene is lost in most members of this clade (Oxelman et al. 2004; Luo et al. 2007), but it occurs both in Escalloniaceae and Apiales. However, clades like Paracryphiales and Bruniales (Lundberg 2001e) have not been sampled for this gene.

Note: Possible apomorphies are in bold. However, the actual level at which many of these features, particularly the more cryptic ones, should be assigned
is unclear. This is partly because many characters show considerable homoplasy, in addition, basic information for all too many is very incomplete, frequently coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. Then there is the not-so-trivial issue of how ancestral states are reconstructed (see above).

Movemenent into the northern hemisphere of Asterales may be linked with the origin of hyperdiverse clades like Asteraceae and Campanulaceae (Beaulieu et al. 2013a), although the basic topology of relationships in Campanulaceae is still unclear and there are suggestions that diversification in Asteraceae began in South America (see that family).

Endress (2011a) thought that the character "monosymmetric flower" in Asterales might be a key innovation, although where it is to be placed on the tree is unclear. Somewhere near the speciose Campanulaceae-Lobelioideae will represent one acquisition of this feature, a position near Asteraceae another. Furthermore, although monosymmetric flowers may occur in most Asteraceae, the capitulum itself is polysymmetric or haplomorphic, and major pollinators behave accordingly (see below under Asteraceae). Endress (2011a) also suggested that a key innovation somewhere in Asterales was tenuinucellate ovules. Unfortunately, corolla and endosperm
development, endothelium presence, not to mention chemistry (for a partial summary, see Grayer et al. 1999), and the like, are unknown in some critical families, so understanding character
evolution is particularly difficult. Absence of apotracheal parenchyma and x = 9 may also be features of Asterales (Lundberg & Bremer 2001; Bremer et al. 2001).

Chemistry, Morphology, etc. The corolla lobes quite often appear to consist of a central portion and marginal "wings" reflecting the induplicate-valvate corolla aestivation of such flowers. For a study of petal vasculature, which shows interesting variation, see Gustafsson (1995); this work needs to be extended. Monosymmetry is often associated
with a slit the length of the corolla, i.e. is the 0:5 type. Variation of ovary position in Asterales is considerable.

Tobe and Morin (1996) summarize embryological knowledge of many members of the order. For some inflorescence morphology, see Philipson (1953), for fructans/inulins, see Meier and Reid (1982), for integument thickness, see Inoue and Tobe (1999), for some inflorescence development, see Harris (1999), and for pollen, see Polevova (2006). For general discussions of variation in the order, see J. Kadereit (2006) and Lundberg (2009).

Phylogeny. For the relationships of Asterales, see the asterid IIclade. Extensive phylogenetic structure in Asterales, although
often with rather weak support, was early apparent (Gustaffson & Bremer 1997; D. Soltis et al. 2000; see also Olmstead
et al. 2000). Subsequent studies improved support for many clades, although there was still a basal polytomy (Kårehed et al. 2000; Lundberg 2001a, b; Kårehed 2002a; especially Bremer et al. 2001; Lundberg & Bremer 2001, 2003). Olmstead et al. (2000) and B. Bremer et al. (2002) suggested a sister group relationship between Campanulaceae and Stylidiaceae (but not Donatia), and the latter authors suggest that Pentaphragmataceae were also associated. Donatia itself is there very weakly linked with Alseuosmiaceae et al., and in some studies it is sister to Abrophyllum
(Carpodetaceae: Gustafsson et al. 1997), not to Stylidiaceae. Stylidiaceae and Donatiaceae are weakly (D. Soltis et al. 2000) or quite strongly (Kårehed et al. 2000; Lundberg 2001; Tank & Donoghue 2010) supported as sister taxa.

There were suggestions that Rousseaceae, Pentaphragmataceae and Campanulaceae were together sister to the other Asterales (Lundberg & Bremer 2003), although the support was not very strong, while Soltis et al. (2007a) found Campanulaceae to be sister to rest of Asterales (1.0 p.p.). Rousseaceae s.l. are often found to be sister to Campanulaceae (Kårehed 2002a; Tank et al. 2007; esp. Tank & Donoghue 2010), but Bell et al. (2010) found Roussea to be sister to the rest of the order. Soltis et al. (2011) found that Pentaphragmataceae were sister to all other Asterales, but with little support - and perhaps because of the pull of some mitochondrial genes.

Relationships between Asteraceae and its immediate relatives also vary somewhat according to the gene studied (A.P.G. 2003 for references). Menyanthaceae did not link with the other three families in the four-gene study of Albach et al. (2001b). Leins and Erbar (2003b) thought that Goodeniaceae were probably sister to Asteraceae, noting i.a. that Barnadesia polyacantha has a bulge beneath the style branch, perhaps homologous with the stylar cup of Goodeniaceae, while Soltis et al. (2007a) found the relationships [[Calyceraceae + Goodeniaceae] Asteraceae]. However, relationships along the spine of Asterales were quite well resolved in a ten chloroplast gene analysis of Tank and Donoghue (2010) and are followed here; Soltis et al. (2011) found a largely similar topology, apart from the position of Pentaphragmataceae and a weakly supported [Phellinaceae [Alseuosmiaceae + Argophyllaceae]] clade.

Previous Relationships. The Asterales here are basically Takhtajan's
(1997) Asteridae, but with the addition of sundry Hydrangeales. Cronquist (1981) also placed many families here in Asterales in the orders placed towards the end of his Asteridae, although some families were also in his Cornales (Rosidae), etc.

Rousseaoideae may be recognised by their whorled, serrate
leaves, scaly buds, and their single, terminal flower with a large calyx that
persists reflexed on the fruit, long-sagittate anthers, and an apparently superior
ovary tapering gradually into a stout, persistent style.

Evolution.Divergence & Distribution. Mauritius is only some 8 m.y.o. - what is the history of Roussea (Lundberg 2001a)?

Pollination Biology.Roussea is pollinated by a gecko (the pollen is embedded in a slimy substance), which may also disperse its seeds (Hansen & Müller 2009).

Chemistry, Morphology, etc.Roussea in particular is poorly known. It has an endodermis in its
petiole, and its seed is drawn as if it
were carunculate (Engler 1930a). Mauritzon (1933) suggested that it might have bitegmic ovules.

Abrophyllum and Cuttsia
both have clusters of small, unlignified cells in the mesophyll that look like
little white raphide bundles (Hils 1985). For a useful summary, see Gustafsson (2006).

For further details of vegetative anatomy of Carpodetoideae,
see Gornall et al. (1998) and Carlquist (2012c), indumentum, see
Al-Shammary and Gornall (1994), floral morphology, see Tobe and Raven (1999),
and seed anatomy, see Takhtajan (2000). For anatomy of Roussea, see Watari (1939) and
Ramamonjiarisoa (1980). For some general
information, see Gustaffson and Bremer (1997) and Koontz et al. (2006: Roussea).

Previous Relationships. Rousseaceae were
previously of uncertain position. Takhtajan (1997) placed Roussea (as Rousseaceae) in
Rosidae-Celastranae-Brexiales, and Carpodetoideae have often been associated with Saxifragaceae
s.l., i.e. the woody Saxifragaceae, thus Takhtajan's Carpodetaceae were members of his heterogeneous Hydrangeales (see also summary in Lundberg 2001a). No members of the family were mentioned by Cronquist (1981), which was perhaps wise.

Age. Bell et al. (2010: Campanuliodeae + Lobelioideae) estimated a crown group age of (67-)56, 53(-41) m.y. for the family, Wikström et al. (2001) suggested an age of (62-)59, 46(-43) m.y., and Knox (2014) and age of around 60 m. years.

1/65. Especially
South Africa, also east Africa (map: from Thulin 1978).

Synonymy: Cyphiaceae A. de Candolle

Campanulaceae are usually herbs that can be recognised by their white latex,
often rather soft, spirally inserted leaves, and flowers with an inferior ovary and secondary
pollen presentation.

Polysymmetric campanulate flowers with sprawling
dehisced anthers at the base inside distinguish Campanuloideae; pollen is held among
hairs along the style, and these hairs later retract. There are often three carpels.

Lobelioideae have monosymmetric flowers in which the
corolla may be split to the base down one side; the anthers (and filaments) are connate and the former at
least initially enclose the stigma. The
style pushes up within the anther tube and exposes the pollen. There are two carpels.

Nemacladoideae are often small herbs that have monosymmetric flowers in which the
filaments are more or less connate and the anthers are free, flipping backwards after they have dehisced. There are two carpels.

Cyphocarpoideae have monosymmetric flowers with a single adaxial lobe that has an apical appendage, the stamens are free from each other but adnate to the corolla, and the inferior ovary, with two carpels, is much elongated; dehiscence of the fruit is through the sides.

Cyphioideae are herbs with tuberous roots that also have monosymmetric flowers in which the
stamens are more or less completely free. There are two carpels.

Evolution.Divergence & Distribution. Knox et al. (2006, no Nemocladoideae or Cyphocarpoideae included) suggested that [Cyphia + Lobelioideae] originated in southern Africa, dispersing quite widely, and with at least two returns to Africa; Antonelli (2009) also suggested that Lobelioideae originated in Africa, and with much subsequent long distance dispersal of the tiny seeds. For ages of various clades within Campanulaceae, see Roquet et al. (2009); ages for deeper nodes in different analyses varied considerably. Crowl et al. (2014) give dates for branching points within Campanuloideae.

The various chromosomal rearrangements studied by Knox (2014 and references) can be integrated with the tree once its topology has settled down.

There has been extensive diversification in the Siphocampylus-Burmeisteria-Centropogon-Lysipomia clade in South America, particularly along the Andes; Lysipomia includes ca 40 rosulate species with small to minute (ca 3 mm across) and sometimes almost polysymmetric flowers growing in the páramo (West & Ayers 2006; Sklenár et al. 2011; see also Knox et al. 2008; Antonelli 2008; Lagomarsino et al. 2014). The Burmeistera clade in particular has had very high speciation rates (e.g. Pennington et al. 2010). Diversification of the whole clade began 18-15 m.y.a., and the 550+ species of the first three genera represent a radiation dating back 12-5 m.y. (Lagomarisono 2014 and references). The pachycaul giant lobelias are derived from herbaceous ancestors (Knox et al. 1993), and giant lobelias from widely separated parts of the globe (Pacific, South America, Africa) may be in the same immediate clade (Antonelli 2009), indeed, some South American taxa may be derived from within the African giant lobelia clade, although the relationships of the giant lobelias and the biogeographic implications of these relationships need more detailed study. Long-distance dispersal is also implicated in the occurrence of Lobelia loochooensis in the Ryukus; it probably came from Australia ca 7,000 km distant (Kokubugata et al. 2012).

Lobelioideae represent a major plant radiation on the Hawaiian islands. Givnish et al. (2006a, 2009a; see also Buss et al. 2001 - seed morphology) note that the some 130 species of Hawaiian Lobelioideae appear to have evolved from a single woody ancestor a mere ca 13 m.y.a. (ages in Antonelli 2009 are slightly older); these ages are older than that of the oldest island, but presumably there was movement from islands that subsequently have sunk. Species there have a variety of growth habits and leaf morphologies, and some species are spiny; herbivory by the now-extinct moa-nalo, a flightless duck as large as a small turkey, is suspected as driving some of this variation. Fleshy fruits have evolved more than once in Hawaii, and pollinators and fruit dispersers also vary considerably (Carlquist 1970; Givnish et al. 1995).

The biogeographic history of Campanuloideae is complex and involves much movement, the area from the Balkans to western Asia being particularly critical in its diversification (Roquet et al. 2009 for details); there are over 100 species of Campanula in Turkey alone. Within Campanuloideae there are several remarkable distribution patterms, although sampling needs to be improved; are Wahlenbergia linifolia (St Helena) and W. berteroi (Juan Fernandez sister taxa (Haberle et al. 2009)? Interestingly, Campanuloideae on Crete seem to be largely remnants of a flora that was on the island when it was originally isolated (Cellinese et al. 2009). Crown Wahlenbergia is (45.3-)29.6(-15.2) m.y. old, stem Wahlenbergia is ca 32 m.y. old (HPD: Prebble 2011); there was little diversification for ca 10 m.y., and W. krebsii, from the Cape, is sister to the other species of the genus sampled.

Pollination Biology & Seed Dispersal. For the evolution of the secondary pollen presentation devices in the family, see Erbar and Leins (1988b) and Leins and Erbar (especially 2003a [Cyphia], 2003b, 2005 [Cyphia], 2006, 2010). However, sampling (inc. Nemocladoideae, some Lobelioideae) is still incomplete, and understanding the evolution of these mechanisms awaits a better supported phylogeny (see below). The protandrous flowers are polysymmetric in bud and the introrse anthers are more or less connivent when they dehisce, pollen then being in a position suitable for secondary pollination, whether entangled with the filament hairs or held immediately above the stigmatic head (e.g. Leins & Erbar 2003b, 2010).

Campanuloideae have brush pollination. Here the pollen is caught in a brush of hairs on the style whence they are removed by the pollinator; in the female phase, the hairs retract so any grains present fall off and selfing is prevented. In Petromarula the stigmatic head is swollen and hairs occur only there (Igersheim 1993a), but otherwise pollination is similar. Phyteuma has coherent corolla lobes although the corolla is open laterally; the style hairs are only partly retractile. The nectar of some Campanuloideae may be brightly colored
and then the filament bases are not persistent; normally they are, and they enclose the nectar. Insect pollination is prevalent, but bird (and lizard) pollination is also known, especially in taxa that grow on islands (Olesen et al. 2012).

In Lobelioideae the pollen is retained in a tube formed by the connate anthers; it is forced out by the stigma before the stigmatic lobes separate, recurve, and finally become receptive. High-altitude species of Burmeistera (Lobelioideae) have both bird and bat pollination (Muchhala 2006); the latter species show character displacement, sympatric taxa differing more in floral morphology than would be expected, so reducing the chances of pollen being deposited on the wrong stigma (Muchhala & Potts 2007). In Centropogon nigricans there seems to have been co-evolution with a remarkably long-tongued bat, Anoura fistulata (Muchhala & Thomson 2009: c.f. Angraecum - Orchidaceae). All told, some 110 species of Andean Lobelioideae may be bat pollinated (Dobat & Peikert-Holle 1985), although the figure in Fleming et al. (2009) is only 20, while L. Lagomarsino (pers. comm.) estimates about 180 species. Extrafloral nectaries are found in Andean "Centropogon" on the outside of the inferior ovary. These occur mostly in species growing at lower altitudes where ants are to be found, and generalist humming birds are usually the visitors; in species at higher altitudes such nectaries were rare and pollination was by sickle-bill humming birds Eutoxeres, although there does not seem to be a connection between nectaries and pollinators (Heliconia [Heliconiaceae] is the nectar resource for Eutoxeres at lower altitudes: Stein 1992). Lobelioideae have radiated extensively on Hawaii, and the flowers of many species of Cyanea and Clermontia (which separated from each other ca 9.7 m.y.a.) are conspicuously curved; pollination of around 125 species on the archipelago is/was by a few species of birds of the extinct and extant Drepanidae and extinct Mohoidae (Carlquist 1970; Lammers & Freeman 1986; Givnish et al. 1995; Pender et al. 2014; T. J. Givnish pers. comm. x.2013). Some species of Clermontia have petaloid sepals, a feature that may have been lost twice (Givnish et al. 2013).

Within Andean Lobelioideae there has been extensive switching betweem fleshy and dry fruits, the result being that Siphocampylus (defined as having capsules) and Centropogon (berries) have turned out to be poly/paraphyletic (Lagomarsino et al. 2014).

Genes & Genomes. For the very extensive rearrangements in the chloroplast genome, including the inverted repeat, see Cosner et al. (1997, 2004), Knox and Palmer (1999: Cyphocarpus,
Nemacladus, etc., not studied, Cyphia was), Haberle et al. (2008a). The chloroplast gene accD (= ORF512, zpfA) has been lost (Doyle et al. 1995 and references; see also Knox 2014). All told, over 125 inversions, most sizable, are known, and Knox (2014) has pieced together the sequence in which some of the larger inversions occurred. Along with these inversions, protein-coding genes, probably from the nucleus, have moved into the chloroplast (Knox 2014). Biparental transmission of plastids has been recorded (Corriveau & Coleman 1988).

Chemistry, Morphology, etc. Details of the major variation patterns in secondary metabolites within the clade need to be established.

Since the pedicel of Lobelia and its relatives is twisted
(resupinate), the flowers appears to have a "normal" orientation with the median petal abaxial, however, this does not usually occur in Lysiopoma (= pseudo-resupinate - Ayers 1997). Some species of Cyphia have a long corolla tube, but the two abaxial petals have slits at their bases; when the flowers are strongly bilabiate, the two abaxial petals are free (Thulin 1978). In Nemacladus (Nemacladoideae) there are sometimes groups of remarkable almost hand-like (in some S.E.M.s) ?nectaries at the bases of two adaxial filaments. It is not known if the style hairs there are retractile. Ostrowskia
(Campanuloideae) has anthers with placentoids and the integument is massive
(Kamelina & Zhinkina 1998).

Some species of Wahlenbergia have an almost superior ovary. Kausik and Subramanyam (1945, 1947), Rosén (1932, 1949) and Subramanyam (1949, 1970) discuss endosperm development; there may be taxonomically interesting variation in cell number of the chalazal endosperm haustorium.

For stem-node anatomy of Campanuloideae, see Col (1904), for flowers of Downingia, see Kaplan (1969 and references), for morphology, see Shulkina et al. (2003), for fruit morphology, see Kolakovsky (1985). Further
general information on the subfamilies was taken from Schönland (1889), Wimmer
(1968) and Lammers (1998 and especially 2006); for pollen see Dunbar (1975a, b), Eddie et al. (2010) and Hong and Pan (2012), Shamrov and Zhinkina (1994) and Shamrov (1998) for ovules, and see Murata (1995), Buss et al. (2001), Cupido et al. (2011 and references) and Koutsovoulou et al. (2013) for seeds, seed coat anatomy/morphology and germination; for the protein bodies in the nuclei, see Bigazzi
(1986) and Haberle (1998), and for the remarkable Nemacladus, see Morin and Ayers (2011).

Phylogeny. The relationships of the major groupings in Campanulaceae were unclear for quite some time. Both the
monophyly and the relationships of the poorly known Cyphiaceae (there are three
subgroups) have been unclear (Lammers 1992), although they are certainly to be included
within a monophyletic Campanulaceae s. l. (see Cosner et al. 1994; Gustafsson
& Bremer 1995; Gustafsson 1996b; Gustafsson et al. 1996). ITS sequence data suggested that one of these groups, Cyphocarpus, was a member of
Lobelioideae (Haberle 1998, Ayers & Haberle 1999) - see also pollen (Dunbar 1975b), but if so, it has several characters in common (parallelisms?) with the Campanuloideae and Nemacladoideae (another element in the old Cyphiaceae). The tree presented by Haberle (1998) suggests the groupings [[Nemacladoideae + Campanuloideae] [Cyphioideae + Lobelioideae]] (Cyphia is the third element), which may imply that the polysymmetric flower of Campanuloideae with the median sepal adaxial (the "normal" condition) is a reversal from a monosymmetric flower with the median sepal abaxial. Lundberg and Bremer (2003) found what was basically a trichotomy of Cyphia, Lobelioideae, and Campanuloideae. Tank and Donoghue (2010) suggested that Cyphia is sister to Campanuloideae and Pseudonemacladus to Lobelioideae; consistent with these relationships is the [[Cyphia + Campanuloideae] clade found by Knox (2014).

General relationships within Campanuloideae are discussed by Eddie et al. (2002, 2003), Cosner et al. (2004), and Olesen et al. (2012). Platycodon, Codonopsis and Cyananthus form a clade and are strongly supported as being sister to all other Campanuloideae (e.g. Cosner et al. 2004; Crowl et al. 2014). Both Campanula and Wahlenbergia are seriously polyphyletic. Most of Wahlenbergia and some other genera form a clade sister to the rest of the subfamily, while within the latter, W. hederacea (now = Hesperocodon hederacea), links rather with Jasione (Haberle et al. 2008b, 2009; Cellinese et al. 2009; Roquet et al. 2008, 2009; Borsch et al. 2009; Prebble et al. 2010; Cupido et al. 2013; Crowl et al. 2014). The terminal Campanula clade can be divided into two main clades, one centred on Campanula s. str. and the other on Rapunculus, while successively basal to these (the relative order is unclear) are the Jasione (mentioned already) and Musschia clades, the latter also including some species of Campanula (e.g. Cupido et al. 2013; Crowl et al. 2014).

Within Lobelioideae, molecular data suggest that Lobelia itself is wildly paraphyletic (Knox & Muasya 2001; Antonelli 2008, 2009; Knox et al. 2008; Lagomarsino et al. 2014). Within South American lobelioids, Centropogon and Siphonocampylus, for example, are not monophyletic (e.g. Lagomarsino et al. 2011, 2014), but there fair support for many relationships in the whole "Siphocampylus"-Burmeisteria-"Centropogon"-Lysipomia clade.

Classification. A.P.G. II (2003) suggested as an option keeping Lobeliaceae separate from Campanulaceae, but the two are best combined in view of their substantial similarities (see A.P.G. III 2009). For a world checklist and bibliography, see Lammers (2007), while Lammers (2011) provided a sectional classification for Lobelia in its current circumscription.

Generic limits need much attention, with a much more broadly delimited Campanula being a reasonable solution to its extensive paraphyly, its segregate genera being based on floral features which are unreliable guides to broad relationships (Haberle et al. 2008b; Roquet et al. 2008). Cupido et al. (2013) outline possible taxonomic solutions to the developing patterns of relationships centred on Wahlenbergia, while Hong and Pan (2012) suggested the generic pulverization of the Codonopsis area. The current classification of Lobelioideae is of little use, and considerable expansion of Lobelia may be a course to take. Thus the monophyletic Burmeisteria is embedded in a clade where Siphocampylus and Centropogon are hopelessly intermingled (most of the infrageneric groups are also para/polyphyletic) , the whole being a clade coming from within a paraphyletic Lobelia (Lagomarsino et al. 2014).

Previous Relationships. Takhtajan (1997) divided Campanulaceae
s.l. into four families; these, plus Pentaphragmataceae and Sphenocleaceae (for the latter, see Solanales),
made up his Campanulanae. Cronquist (1989) had taken the opposite tack, recognizing a broadly circumscribed Campanulaceae that included the four families just mentioned, plus some other families that are elsewhere in Asterales, in his Campanulales.

Pentaphragmataceae are distinctive plants. They are rather fleshy herbs with two-ranked, asymmetric, usually serrate leaf blades and scorpioid cymes
with sessile flowers. The flowers are
radially symmetric and have relatively large, conspicuous sepals, extrorse anthers and an inferior ovary; there are nectariferous
pockets between the septae that join the hypanthium to the ovary. The fruit is baccate.

Chemistry, Morphology, etc. The family is very poorly
known. The micropylar haustorium is single-celled (Kapil & Vijayaraghavan 1965).

For general information, see Lammers (2006), for wood anatomy, see Carlquist
(1997b), for the flower, see Vogel (1998b), and for pollen, see Dunbar (1978).

Phylogeny. There is a possible grouping [Alseuosmiaceae [Phellinaceae + Argophyllaceae]] or [Alseuosmiaceae [Stylidiaceae [Phellinaceae + Argophyllaceae]]] (e.g. Kårehed et al. 2000; Lundberg & Bremer 2001 respectively), and although jacknife support for the
position of Alseuosmiaceae is not very strong, the posterior probability for the first grouping is 1.0 (Kårehed 2002a).

Alseuosmiaceae may be recognised even vegetatively by
their spiral, serrate leaves with axillary tufts of uniseriate hairs. Their flowers have an inferior ovary
and usually a tubular corolla, and the corolla lobes have fringed or erose winged
margins. The fruit is baccate and
contains rather small seeds.

Evolution.Plant-Animal Interactions.Platyspermation and other Alseuosmiaceae on New Caledonia commonly have galled fruits or flowers.

Chemistry, Morphology, etc. Ellagitannins are reported from Alseuosmia (Kårehed 2006 for references); this should be checked. Most Alseuosmiaceae have rayless wood, living
mature fibres with stored starch (Dickison 1986b), and the stem has an endodermis. Uniseriate hairs in Platyspermation are not restricted to the leaf axils, although they are particularly dense there, rather, they cover the whole plant. Their persistent, reddish bases look rather like glands, hence, perhaps, the past inclusion of the genus in Rutaceae.

There are tanniniferous cells in the flower. The margins of the corolla lobes of Platyspermation have narrow flanges and papillae; the corolla is only shortly tubular, the lobes being rather spreading (buzz pollination?). The pollen of Alseuosmia linariifolia is described as being tricolpate, with an ectexine made up of a thick, tubercular tectum and massive, spherical columellae (Polevova 2006); whether this can be generalised to the family is unclear (see also Kårehed 2006).

Some details of vegetative anatomy are
taken from Paliwal and Srivastava (1969), Dickison (1989a) and Gornall et al. (1998), and of testa anatomy from
Nemirovich-Danchencko and Lobova (1998) and Takhtajan (2000). The embryology is poorly known. See Kårehed (2006) for general information.

Phylogeny.Platyspermation is strongly associated with other Alseuosmiaceae (Lundberg & Bremer 2001), and may be sister to the rest of the family (Tank & Donoghue 2010).

Classification. Generic limits are in some dispute (Tirel 1996).

Previous Relationships. Genera now included in Alseuosmiaceae have previously been placed in
Caprifoliaceae, Rubiaceae, Rutaceae, Ericaceae, Epacridaceae, etc. Although the family was recognized by Takhtajan (1997), it was included in his
Hydrangeales.

Chemistry, Morphology, etc. The
guard cells are huge, with inner and outer stomatal ledges. The ovules are reported as being hemitropous to campylotropous, but Phelline is embryologically poorly known.

See Baas (1975) for wood anatomy
(it appeared to be extremely primitive), Lobreau-Callen (1977) for pollen, and Kårehed et al. (2000) and Barriera et al. (2006) for much
additional information.

Previous Relationships. Takhtajan (1997) placed the family in Icacinales,
describing the leaves as being mostly estipulate, while Cronquist (1981) placed it in his Aquifoliaceae (adjacent to Icacinaceae), both were in his Celastrales.

Stylidiaceae are usually rosette or tussock herbs with sessile or obscurely petiolate leaves. Stylidioideae have very
distinctive semi-resupinate, often split-monosymmetric flowers. The two stamens are usually adnate to the style, and the whole complex is often
sensitive,
moving when brushed by pollinator (hence the name, trigger plants). Donatioideae are small-leaved tussock plants that can be
recognised by their solitary, polysymmetric, terminal flowers that have
a variable number of free sepals and petals and two to three stamens with extrorse
anthers.

Evolution.Pollination Biology. In many Stylidioideae (Stylidium s.l.) the two stamens are adnate to the style, the
extrorse anthers being borne near the stigma. The whole complex (a gynostemium), sometimes called a column, is often
sensitive,
moving rapidly when brushed by the pollinator; in some species the column is hinged. For the literature on stylar movement, see Findlay and Findlay (1989). In Levenhookia, however, the gynostemium is held under tension in the hooded labellum, flipping only when the latter is disturbed. Armbruster et al. (1994) found that species of Stylidium in the Perth area were pollinated mostly by solitary bees and bombyliid flies, species in the same locality differing in both corolla tube and column lengths; even the pollen varied in colour.

Ecology & Physiology. There are suggestions that Stylidium may be carnivorous. Insects are trapped by the glandular hairs, which also show yeast-extract stimulated protease acivity; the plants grow in acid, nutrient-poor soil like other carnivorous plants. However, uptake of nutrients by the plant from the insects has yet to be demonstrated (Darnowski et al. 2006).

Chemistry, Morphology, etc. In Stylidioideae the cambium
may develop beneath the endodermis; xylem, and sometimes also phloem, is produced
towards the inside, and at most cork to the outside (Carlquist 1981a, 2013). In older stems of Donatia the cortex is very
thick, and the vascular tissue forms a narrow cylinder in the center (Chandler
1911).

The fertile stamens are the adaxial pair. Carolin (1960b) drew the anthers of Donatia as being introrse. Ronse de Craene (2010) described the flower of Stylidium graminifolium as being obliquely monosymmetric at maturity, and the corolla and parts inside are illustrated as having rotated ca 60o relative to the calyx, and there is a single abaxial (adaxial in the text) nectary. See also Erbar (1992) for floral development, which needs more study in the family as a whole. The pollen of at least some Stylidiaceae has a very distinctive inner ectexine that lacks columellae but is permeated by numerous sinuous channels (Polevova 2006). Monocotyly is reported to be quite common in Stylidioideae (Carlquist 1981b).

For
anatomical differences between Donatioideae and Stylidioideae, see Repson
(1953). The proembryo in Donatia
is ovate, the suspensor is made up of short cells, but in Stylidioideae it is long, and is made up of cells that are longer than broad (Philipson &
Philipson 1973), as in other Asterales (Tobe & Morin 1997). The leaves of Donatia are very small, and
their venation is acrodromous.

For general information, see Carolin (2006), Carlquist and Lowrie (1989: Stylidioideae), Australian Plants 27(215). 2013, and Glenny (2009: Forstera); some anatomical details can be
found in Thouvenin (1890), for embryology, see Rosén (1935) and Subramanyam (1951a), for placentation, see Carolin (1960a), for protein bodies, see Thaler (1966), and for the testa anatomy of Stylidium, see Tobe and Morin
(1996).

Phylogeny.Donatia is sister to the rest of the family, but there is some uncertainty over further relationships. Laurent et al. (1999) found that Forstera s.l. was sister to the rest of the family in combined molecular and morphological analyses, in a rbcL + ndhF analysis it was grouped with Levenhookia, but both positions had only weak support; Wagstaff and Wege retrieved the former topology in an analysis based on variation in ITS and rbcL. In gross floral morphology Donatia and Forstera are similar, both having basically radially symmetrical flowers that are whitish in colour.

Previous Relationships. Stylidiaceae have been treated as two families in Stylidiales (Takhtajan 1997)
or merged in one family (Philipson & Philipson 1973). A.P.G. II suggested as an option keeping Donatiaceae and Stylidiaceae separate, although the two can reasonably be combined (e.g. Lundberg & Bremer 2003; A.P.G. III 2009).

Evolution.Divergence & Distribution. Pollen of all families of this clade - and of some subfamilies of Asteraceae - had differentiated by the Oligocene, and has been found in many places that are fragments of the Gondwanan continent (Barreda et al. 2010a).

The separate vascular bundles in the stem may be connected with the herbaceous habit that is so common here. Presence of sclereids, bicellular pollen, and multi-nucleate tapetal cells may also be synapomorphies for this clade (Lundberg 2009, q.v. for more possible synapomorphies).

Chemistry, Morphology, etc. The androecium has spiral initiation in some
Menyanthaceae, Asteraceae and Goodeniaceae - and also Araliaceae (Erbar 1997). For other characters common to this clade, see Anderberg et al. (2006, c.f. vessel perforation plates). Vegetatively and florally Menyanthaceae are rather different from many other Asterales, however, both Menyanthaceae and Goodeniaceae have corolla lobes with marginal wings, here placed as apomorphies for both groups (or they could be a synapomorphy for the whole clade, being lost later). For inflorescence morphology and evolution, see Pozner et al. (2012).

Menyanthaceae are aquatic plants with more or less orbicular or
palmately-compound leaves with broad bases and quite large usually
heterostylous polysymmetric flowers. The petals are connate, the corolla lobes have disinctive marginal
wings or fringes and the ovary is superior.

Age. An estimate of the age of crown-group Menyanthaceae is (58-)54, 51(-47) m.y. (Wikström et al. 2001) and another is (60-)47, 44(-31) m.y. (Bell et al. 2010).

For fossil pollen, see Barreda et al. (2010a).

Evolution.Divergence & Distribution. Has the superior ovary of Menyanthaceae with its parietal placentation been derived from a more or less inferior ovary with axile placentation (c.f. Pittosporaceae - Apiales)?

Eichler (1878) drew the flower of Menyanthes with an oblique plane of symmetry. The vascular anatomy of
the flower implies a basic monosymmetry - the lateral corolla traces are fused (Wood
& Weaver 1982). There are sometimes fringed scales, "staminodes", on the corolla tube alternating with the stamens. Johri et al. (1992) suggested that the endosperm stores starch.

For embryology, see Stolt (1921) and Maheswari Devi (1963), for seed morphology, see Chuang and Ornduff (1992), for floral development, see Erbar (1997), for the missing rpl2 intron, see Downie et al. (1991b), and for general information, see G. Kadereit (2006).

Phylogeny. Relationships within Menyanthaceae have been clarified by Tippery et al. (2006, 2008) and Tippery and Les (2008); [Menyanthes + Nephrophyllidium] are sister to the rest of the family while Villarsia is very much paraphyletic. Many relationships within Nymphoides are as yet poorly supported; whether or not the flowers are heterostylous seems a very labile character (Tippery & Les 2011).

Previous Relationships. The iridoids of Menyanthaceae differ
chemically from those of Gentianaceae, in which Menyanthaceae used
to be included, although placentation, etc., are similar. Menyanthaceae were placed in Solanales by Cronquist (1981). Branched sclereids and air
canals are similarities between Menyanthaceae and Nymphaeaceae, but both are aquatics.

Evolution.Divergence & Distribution. Although secondary pollen presentation occurs throughout this clade, there is considerable variation in the details of how it is done (Leins & Erbar 2003b, 2006 and references).

Study of early capitulum development in Arnaldoa macbrideana (Asteraceae-Barnadesioideae) suggests that the capitulum there is built up of partial inflorescences with cymose branching, so perhaps linking the racemose heads of Asteraceae with the apparently rather different inflorescences of many Calyceraceae and Goodeniaceae (Leins & Erbar 2003b, see their polytelic thyrses). Acicarpha is the only Calyceraceae with n = 8, and it is also the only member of that family with a possibly plesiomorphic condensed spicate inflorescence (DeVore 1994). For a comparison of the pollen of the three families, see DeVore et al. (2007), for chromosome numbers see Semple and Watanabe (2009).

Ca 8/330: Goodenia (190), Scaevola (100). Throughout Australia, to New Zealand, Chile and China; Scaevola pantropical, with the coastal S. taccada in the E. and C. Pacific and Indian Oceans and S. plumieri in the W. Indian, E. Pacific and the Atlantic oceans (from van Balgooy 1975). [Photo - Flower.]

Age. The age of crown-group Goodenieae is estimated to be 48-)37.1(-27) m.y.a. (Jabaily et al. 2014).

Synonymy: Scaevolaceae
Lindley

Goodeniaceae are more or less herbaceous plants with
spiral leaves and monosymmetric flowers. The corolla is often divided to the base adaxially, the lobes have
marginal wings that may make each lobe look trilobed, and the style is curved, with
an apical cup and (in older flowers) a bilobed stigma in the
middle. The anthers dehisce in bud and
the ovary is nearly always inferior.

Jabaily et al. (2014) discuss ideas about the origin (place equivocal) and time (quite a spread of ages) for the origin of Goodeniaceae. Some diversification in Scaevola may be associated with the aridification of the Nullarbor Plain some 14-13 m.y.a. separating eastern and western clades (Crisp & Cook 2007).

Pollination Biology & Seed Dispersal. For the diversity of pollen presentation devices in the family, see Erbar and Leins (1988) and Leins and Erbar (2003b, 2010). In nearly all species the pollen is initially enclosed by the indusium, and is pushed out by the elongating style; the pollen of Brunonia, with its capitate inflorescence and flowers with straight styles, is held in a brush formed by stylar hairs.

The seeds of many Goodenieae in particular are myrmecochorous (Lengyel et al. 2009, 2010), others have circumferential wings that are mucilaginous when wetted (Jabaily et al. 2012).

Chemistry, Morphology, etc. The cortical bundles in the stem sometimes reported for the family are leaf traces.

The lateral veins of the corolla of Brunonia unite in the receptacle (Erbar 1997). The pollen is distinctive;
endoapertures are bordered (with orae) and lalongate and the spines are rounded
(Gustaffson et al. 1997).

See also Subramanyam (1950a) for embryology, Carolin
(1978, 2006) for general variation and morphology, Carolin (1966) for seeds and fruits, and Erbar and Leins (1988) and Cave et al. (2010) for the floral development of Brunonia.

Phylogeny. Relationships within Goodeniaceae are becoming fairly well understood (Gustafsson 1996a; Gustafsson et al. 1996; Jabaily et al. 2010, esp. 2012). There are two main clades, one includes Lechenaultia and allies, and the other Scaevola, a paraphyletic Goodenia, and allies. Within the latter clade Brunonia is sister to the rest.

Classification. For a general account of most of the family, see Fl. Austral. vol. 35 (1992). Generic limits around Goodenia are difficult and the limits of the genus may have to be severely circumscribed (Jabaily et al. 2012). Most of the numerous features in which Brunonia differs from other Goodeniaceae are autapomorphies (see above). Some of these features might seem to suggest relationships with Asteraceae (Gustafsson 1996a), but clearly the exclusion of Brunonia, as by Cronquist (1981) would make Goodeniaceae paraphyletic (Jabaily et al. 2012).

Evolution.Divergence & Distribution. Both Calyceraceae and Asteraceae-Barnadesioideae, sister to the rest of Asteraceae, are South American, and this whole clade may have originated there.

For other similarities or possible synapomorphies, see DeVore (1994) and Lundberg and Bremer (2001, 2003), these include libriform fibres with simple pits and vasicentric parenchyma. Pesacreta et al. (1994) suggest similarities in the micromorphology of the filaments and connective bases between at least some members of Calyceraceae and Asteraceae.

Placement of characters like pollen with intercolpar depressions on the tree is difficult to ascertain (see also DeVore 1994; DeVore et al. 2000). Indeed, DeVore and Skvarla (2008) suggest that pollen characters thought to suggest a relationship between the two families are different in detail and are therefore not homologous, similarly, although both families have but a single ovule, the position and orientation of the ovule is such that the single ovule condition may well have been derived independently. Finally, the inflorescence, although superficially similar in the two families, may be cymose in Calyceraceae and racemose in Asteraceae.

Understanding more about the expression of CYC (Cycloidea) genes in this clade - and indeed in Asterales as a whole - is likely to be interesting. Chapman et al. (2012) note that Acicarpha (Columelliaceae) has only CYC2a, genes, Dasyphyllum (Barnadesioideae, flowers polysymmetric again) also has CYC2c genes, while the other Asteraceae studied all had CYC2b, d and e genes as well. Members of the CYC2c gene family are involved in the monosymmetric phenotype, where corolla formation on the adaxial side of the flower is more or less reduced.

Chemistry, Morphology, etc. Calyceraceae
and Barnadesioideae have a similar simple flavonoid profile. For inflorescence morphology, see Pozner et al. (2012).

Previous Relationships. Both Cronquist (1981) and Takhtajan (1997) placed the two families in separate, if adjacent, orders.

Calyceraceae are herbs that may be recognised by their
capitate inflorescences the flowers of which often open centripetally. The flowers are small and polysymmetric, the
sepals are spines or are thick and aerenchymatous, the outer layer of the
tubular corolla is photosynthetic, the stamens are free, and the ovary is
inferior.

Chemistry, Morphology, etc. Cronquist (1981) described the flowers as being sometimes "slightly irregular"; they are commonly polysymmetric. There may be five carpels (e.g. Erbar 1993). The integument is described as being "thick" and the outer cell layers contain chloroplasts (Dahlgren 1915).

Some general information is taken from
Hansen (1992: especially useful), DeVore (1994), DeVore and Stuessy (1995), and Hellwig (2006), some details of morphology come from Pontiroli (1963), embryology from Dahlgren (1915: one species), floral development from Erbar (1993: one species, and very odd development at that), inflorescence development from Harris (1999: two species) and cytology from Benko-Iseppon and Morawetz (2000b: one species).

Age. Crown-group Asteraceae are dated to some 42-36 m.y. (K. J. Kim et al. 2005), (52-)43, 40(-31) m.y. (Bell et al. 2010), or (44-)41, 40(-37) m.y. (Wikström et al. 2001); other suggested ages are similar (Funk et al. 2009c for a summary; see also Torices 2010). However, Beaulieu et al. (2013a: 95% HPD) estimated a somewhat older crown-group age of (52-)49(-48) m.y., ages in Swenson et al. (2012) range mostly from (71.1-)52.6, 47.4(-45.4) m.y. (see also Jabaily et al. 2014 for similar estimates), while 47.6-47.3 m.y.a. is the age in Funk et cl. (2014). (Heads [2012] thought that the mostly Antipodean Abrotanella, basal Senecioneae, diverged from the rest of the tribe in the Jurassic or Early Cretaceous, which would thus imply an age for Asteraceae as a whole of around 1,500,000,000 years, or about a third of that age, depending on which vicariance dating you elect to pick [Swenson et al. 2012].)

Asteraceae are very variable vegetatively, but may be
recognised by their capitulate and involucrate inflorescences in which the numerous
small flowers open first on the outside and are only sometimes subtended by
bracts. The anthers are usually fused and
form a tube through which the style extends before the two stigmatic lobes separate
and become recurved. The rather small, single-seeded fruits usually have a plumose pappus (the much-modified calyx), and are frequently dispersed by wind. Sesquiterpene lactones give the bitter taste of many Asteraceae, e.g. Cnicus benedictus.

Evolution.Divergence & Distribution. Papers in Funk et al. (2009a) summarize biogeography, clade ages, and much more for each tribe. For fossil pollen, see Barreda et al. (2010a), and depending on the identification of the grains, the first seven subfamilies in the sequence above may all have diverged by the late Eocene about 34 m.y.a.; other suggestions are of an (early) Oligocene radiation of the subfamilies (K.-J. Kim et al. 2005; Funk et al. 2005; Barker et al. 2008; Torices 2010). See also Funk et al. (2014) for dates.

Diversification of Asteraceae probably began in southern South America - the six basal clades are concentrated in South America - movement to Africa being by way of islands on what are now the Rio Grande Rise and the Walvis Ridge. It has been suggested that Asteraceae making this move may have evolved features common in island plants (Carlquist 1974) like more or less shrubby or tree-like growth forms (Katinas et al. 2013), although a woody/shrubby habit is also common in several of these South American clades (Panero et al. 2104). Many clades then arose in the course of a subsequent radiation from Africa (Panero & Funk 2008; Funk 2009; Funk et al. 2009c; see also Kim & Jansen 1995). Thus both South America and Africa are central to our understanding of the early spread and diversification of the family.

At 2,400+ species the speciose Cardueae makes up most of Carduoideae; the more basal branches are very species-poor. The tribe is now primarily Mediterranean, and Barres et al. (2013) discuss its history in some detail. Much of the diversification in the Mediterranean-Central Asian Carduoideae-Centaureinae may be Plio-Pleistocene transition and younger (Hellwig 2004). Much diversification of the high-altitude species of Saussurea may have occurred in the context of the uplift of the Qinghai-Tibetan plateau within the last 14 m.y. (Y.-J. Wang et al. 2009), J.-Q. Liu et al. (2006) advance a similar explanation for the diversification of the 200+ species of the largely unresolved Ligularia–Cremanthodium–Parasenecio clade, and there have been other radiations of Asteraceae in this area (J. W. Zhang et al. 2011 and references).

Bell et al. (2012b) suggest that there has been rapid diversification of Tragopogon et al. (Cichorioideae) in Eurasia, probably ca 2.6 m.y.a. or at least within the last 5.4 m. years. The closest relatives of the Hawaiian endemic Hesperomannia are from Africa (Kim et al. 1998, see also Keeley & Funk 2011); divergence from Hesperomannia from Vernonia has been dated at around 26-17 m.y.a., old for a Hawaiian endemic (Kim et al. 1998; Keeley & Funk 2011 for a list).

Crown Gnaphalieae (Asteroideae) can be dated to (52.3-)34.5(-20.6) m.y.a., diversification probably beginning in southern Africa, with various subsequent dispersals including one that resulted in the some 550 species of the Australasian part of the tribe (stem age [22.1-]15.6[9.1] m.y.; crown age [20.6-]14.6[-8.3] m.y. - see Bergh & Linder 2009). Within Senecio s. str. there has been intercontinental dispersal between areas with Mediterranean and desert climates (Coleman 2003). Riggins and Siegler (2012 and references) discuss the biogeography of Artemisia, Eurasian in origin; migration here has also been extensive, while Strijk et al. (2012) look at the dispersal of a polyphyletic Psiadia to Madagascar and thence to the Mascarenes.

The iconic stout-stemmed (pachycaul) giant senecios (Dendrosenecio) of the African mountains are all closely related and not immediately related to Senecio s. str. (Knox & Palmer 1995a, b; Pelser et al. 2007). Espeletia is a characteristic genus of the Andean páramo, and it, too, is secondarily woody and often more or less pachycaul, and other Asteraceae are to be found in this habitat (Sklenár et al. 2011 and references). As Small (1919: p. 142) noted of the ability of Asteraceae to colonize places like Krakatau, "The Compositae, indeed, seem to have been formed with the mountains by the mountains for the mountains.".

For possible apomorphic characters for the family and its major clades, see e.g. Hansen (1991), Jansen et al. (1991), Bremer (1994), Leins and Erbar (2000), Erbar and Leins (2000), Funk et al. (2009c), Roque and Funk (2013) and Panero et al. (2014). It is hardly surprising, given the size and ubiquity of the family, that there should be speculation about what has caused its diversification, whether the development of the capitulum itself with its high seed set, the storage of carbohydrates as unbranched-chain fructans, the diversity of secondary metabolites produced, or some other reason (see also Funk et al. 2009c). Burleigh et al. (2006) suggest that by some measures Asteraceae do show a notable shift (increase) in morphological complexity. More attention should be paid to the significance of pollination of Asteraceae by oligolectic bees (see below). There was a palaeopolyploidy event involving most or all of the family, and Schranz et al. (2012) suggested that there was a lag time between the this duplication event and subsequent diversification of the family, which they thought might be causally linked.

There are perhaps parallels with Poaceae, which also have large-scale genome duplications and store carbohydrates as fructans - diversification may be best explained by focusing on particular clades in the family rather than treating the family as a unit (Schranz et al. 2012; see also, perhaps, rate shifts in S. A. Smith et al. 2011). Indeed, although Asteraceae alone contain about 8% of eudicot species, within Asteraceae, Asteroideae, the equal-youngest subfamily, include over 16,000 species, some two thirds of the family; most of the twelve clades basal to Asteroideae have relatively few to very few species (the sister clade of Asteroideae has seven species), although Cichorioideae have ca 3,600 species and Carduoideae ca 2,800 species. Net evolutionary rates within the family are highly heterogeneous (S. A. Smith et al. 2011).

Ecology & Physiology. Growth form is notably labile in Asteraceae, and many taxa are more or less woody-herbaceous intermediates (Beaulieu et al. 2013b: the character there = woody vs herbaceous, but c.f. e.g. Carlquist 2013). Woody Asteraceae are common on ecological/actual islands, where the evolution of woodiness is common (e.g. Carlquist 1974). Climbing Asteraceae are prominent in montane forests of South America (Gentry 1991).

The storage of carbohydrates as unbranched-chain fructans may contribute to the ability of Asteraceae to live in the rather dry conditions that many of them prefer (John 1996).

Flaveria (Asteroidae) has some species with C4 photosynthesis, some with C3, and some intermediate with C2 photosynthesis; details of the metabolic changes involved are quite well understood, and there have been several of these shifts (Bläsing et al. 2000; McKown & Dengler 2009; Ludwig 2011a and references, b, c; Gowik et al. 2011; Schulze et al. 2013; T. Sage et al. 2013; Christin & Osborne 2014); there are also proto-Kranz species (R. Sage et al. 2014). Christin et al. (2011b) suggest a number of dates for C4 origins here, and all are less than 4 m.y.a.; the repeated changes in photosynthetic mechanism may reflect an underlying "predisposition" (McKown et al. 2005), as has been suggested for the evolution of C4 photosynthesis in other groups.

Pollination Biology & Seed Dispersal.

Pollination Biology.

The capitulum can become reduced to a
single flower, the single-flowered capitulae may then reaggregate into a supercapitulum, and there may occasionally be yet another round of aggregation (Claßssen-Bockhoff 1996b for details; Harris 1994 [Caenozoic capitulae], 1999; Leins & Gemmeke 1979; Katinas et al. 2008a). Whether a normal capitulum or some kind of supercapitulum, the whole inflorescence functions as a flower in terms of attracting pollinators, and some genes whose expression is normally restricted to individual flowers may be more widely expressed in the capitulum as a whole, as well as in vegetative shoots (Ma et al. 2008). Disc flowers are quite often polysymmetric and the ray flowers monosymmetric. This monosymmetry seems to be caused by the CYC2c gene family, different members of which have been independently coopted by different taxa in Asteraceae causing cell growth in the adaxial part of the corolla to be reduced, hence the ray phenotype. Interestingly, Dasyphyllum (Barnadesioideae) lacks members of the CYC2b, d and e gene families common elsewhere in Asteraceae, while Acicarpha (Columelliaceae) also lacks CYC2c genes (Chapman et al. 2010).

In general, insect - especially bee - pollination occurs throughout the family, for bird and wind pollination, see below. Pollinators of Asteraceae might seem not to be very selective, since the frequent and diverse insect visitors so obvious on a capitulum of any size trample around on top and appear to pollinate indiscriminately as they go, but this may not be quite true. Effective pollination is commonly carried out by a variety of broadly oligolectic small and often solitary bees belonging to Andrenidae (not in Australia) and Colletidae. These form complex and partly learned associations with individual species of Asteraceae; both bees and Asteraceae are common in drier areas (Linsley 1958; Moldenke 1979b; Lane 1996; Müller & Kuhlmann 2008; Kuhlmann & Eardley 2012). Moldenke (1979b) estimated that in North America about 525 bee species, well over one third of the total number of oligolectic bees, were restricted to Asteraceae. Thus within north temperate Colletes (plasterer bees) a few species specialize on Asteroideae, the other species rarely visiting them; specialization on flowers of Asteraceae has evolved three or four times there (Müller & Kuhlmann 2008). Pollination in Helianthus (sunflowers) has been particularly well studied, and several species of oligolectic bees may visit the one species of sunflower. Thus 39 species of oligolectic bees (mostly Andrenidae and Anthophoridae) and 22 species of polylectic bes visited 21 species of Helianthus regularly for pollen, and although at most few were obligately associated, the majority worked other Asteraceae. All told, 284 species of bees visited sunflowers for pollen, 128 species for nectar (Hurd et al. 1980; also Minckley et al. 1994). Similarly, Schemske (1983) noted that 11-20 species of bees, and over twice as many species of insects in general (in both cases, sometimes many more), commonly visit a single species of Asteraceae.

Interestingly, pollen of Asteraceae-Asteroideae and -Cichorioideae, at least, may be unsuitable for many bees. It may lack essential amino acids, have generally lower amino acid and protein concentrations than other pollen, or contain harmful secondary metabolites (Waser et al. 1996; Müller & Kuhlmann 2008; Goulson 2010; Sedivy et al. 2011). Consequently, some bees actively avoid collecting pollen from composites, thus female bumble bees may get covered in pollen as they collect nectar, yet they do not transfer that pollen to their corbiculae (Neff & Simpson 1990; Goulson 2010), although this would not stop them being effective pollinators (and might even enhance their effectiveness - dirty bees pollinate more efficiently).

Some Asteraceae have a pump
(Nüdelspritze) mechanism of secondary pollen presentation, others have a
brush mechanism (see Leins & Erbar 2003b for a possibly evolutionary sequence of pollen presentation devices). Within Barnadesioideae, sister to the rest of the family, secondary pollen presentation is by simple deposition on the style/stigma (as in at least some Calyceraceae) or by an unspecialised type of brush mechanism (e.g. Erbar & Leins 2000; Leins & Erbar 2006). A number of Carduoideae, especially Centaurineae, have touch-sensitive stamens. The filaments contract when the flowers are touched by the pollinator, and the pollen is then forced out of the anther tube; this pollination mechanism appears to have arisen more than once, and is associated with short and sticky, not long and dry, stigmas and smooth, not spiny, pollen (López-Vinyallonga et al. 2009).

Barreda et al. (2010b, 2012) suggested that the flowers Raiguenrayun, from the Middle Eocene of Patagonia ca 47.5 m.y.a., might be pollinated by birds, but humming birds, the iconic bird pollinators of the New World, are known only from Europe at that time (e.g. Mayr 2004), so bird pollination is unlikely (see also Panero et al. 2014). Bird pollination is rather uncommon in Asteraceae (Cronk & Ojeda 2008).

In wind-pollinated Asteroideae the heads have either staminate or carpellate flowers. In male heads the anthers are free and the capitulae are often pendulous, and the pollen grains have lost their spines. Since the carpellate heads may have only a single flower, the end result is a breeding system very much like that of other wind-pollinated plants like Fagales - monoecy, aggregated pollen-producing units, and female reproductive units that produce a single-seeded fruit. For the phylogeny, genome evolution, etc., of the wind-pollinated Artemisia, with its multiple invasions of the Arctic (polyploidy is apparently not involved), see Vallès and Garnatje (2005), Sanz et al. (2008) and Trach et al. (2008).

There is a great diversity of breeding
systems in the family (e.g. Burtt 1961, 1977a), and the evolution of different flower types in Inuleae (Asteroideae) has been examined by Torices and Anderberg (2009). Although protandry is very common, when there are different flower types, interfloral protogyny predominates (Bertin & Newman 1993). Apomixis is quite common, as in Hieracium and Taraxacum (see T. absurdum!), both in Cichorioideae, and Antennaria (Asteroideae). Dioecy has evolved from monoecy and back again in Leptinella (Cotula s.l.: Himmelreich et al. 2012).

Seed Dispersal.

Most fruits are crowned by a plumose pappus, a highly modified calyx (Yu et al. 1999: confirmation at the level of gene expression; Mukherjee & Harris 1995 and Nordenstam 2008 for variation). The hairs that make the pappus up are themselves sometimes hairy, and wind dispersal is very common. However, a number of taxa are dispersed by animals, whether by hooked fruits (Bidens), or hooks on the inflorescence (Arctium), or by myrmecochory, the fruits having some sort of elaiosome, as in Centaurea (Carduoideae) and Osteospermum (Asteroideae: Lengyel et al. 2009).

Plant-Animal Interactions. The flowers of some Senecioneae and Eupatorieae (Asteroideae) are visted by male Danainae and Ithomiinae (butterflies) and Arctiinae and Ctenuchidae (moths) and of larvae of Arctiinae because the pyrrolizidine alkaloids they contain form the basis of their pheromones, or of compounds that other organisms find distasteful (see also Crotalaria, Heliotropaceae, and Apocynaceae: Edgar et al. 1974; Fiske 1975; Ackery & Vane-Wright 1984; Brown 1987; Weller et al. 1999; Anke et al. 2004); for the phylogeny of Arctiinae and the evolution of pharmacophagy there, see Zaspel et al. (2014). The tribes are unrelated, so there has been parallel evolution of these alkaloids within Asteroideae (e.g. Reimann et al. 2004). Pyrrolizidine alkaloids and pentacyclic triterpene saponins obtained from Asteraceae and variously modified are also found in the secretions of the defensive glands of some Chrysolina and Platyophora beetles (Chrysomelidae); both genera are very speciose (Pasteels et al. 2001; Termonia et al. 2002; Hartmann et al. 2003). Pyrrolizidine alkaloids protect the plants that have them against some herbivores, although individual alkaloids in Senecio section Jacobaea are readily and seemingly randomly gained and lost during evolution by the switching on or off of the genes involved in their synthesis, and there is also much variation in the amount of individual alkaloids (Pelser et al. 2005). Sesquiterpene lactones from Asteraceae are also sequestered by insects (e.g. Pasteels et al. 2001).

Larvae of Nymphalidae-Melitaeini butterflies are common on Asteraceae, as well as on Lamiales, from whence they probably moved less than 50 m.y.a. (Wahlberg 2001; Nylin & Wahlberg 2008; Nylin et al. 2012), a move perhaps associated with an increase in their diversification rate (Fordyce 2010). Caterpillars in a clade of Nymphalidae-Heliconiinae-Acraeini utilise primarily Andean Asteraceae, probably switching from host plants in the Passifloraceae area (Silva-Brandão et al. 2008), but in this case without a change in diversification rates (Fordyce 2010). Agromyzid dipteran leaf miners have diversified in north temperate Asteraceae; these insects prefer plants with noxious secondary metabolites (Winkler et al. 2009).

Within Carduoideae the stout root stocks and large flower heads in particular are resources for the numerous herbivorous insects that specialize on this clade. More than fifty genera of specialized thistle insects, including representatives of Zygaenidae, Tortricidae, Pterolonchidae (all Lepidoptera), Curculionidae (Coleoptera), Tephritidae (Diptera), Tingitidae (Hemiptera) and Cynipidae (Hymenoptera), are found on Carduoideae of the west Palaearctic region, although their numbers are not great considering the diversity of Carduoideae there (Zwölfer 1988; Csoka et al. 2005; Brändle et al. 2005 and literature). All these herbivores are particularly abundant in the Mediterranean region, which is perhaps where Carduoideae evolved (Zwölfer 1988). Introduced insects including a weevil and the tephritid Urophora are often very effective biological control agents of introduced Carduoideae in North America and other parts of the world (Redfern 2011). Tephritid flies are particularly noteworthy on Carduoideae, either eating fruits, exudates they induce from the plant, or forming galls in the stem or inflorescence (Korneyev et al. 2005, esp. Urophora; Redfern 2011). They are also common on other species of the family pretty much world-wide (e.g. Prado & Lewinsohn 2004; Norrbom et al. 2010), and tephritid-induced ball galls on the stems of Solidago (Asteroideae) growing in the prairies of North America are conspicuous in the late summer (Abrahamson & Weis 1997). Interestingly, Prado and Lewinsohn (2004) found that related species of Asteraceae in the Espinhaço mountains or Minais Gerais, Brazil, tended to support a similar tephritid fauna which, however, might not be made up of immediately-related taxa.

Bacterial/Fungal Associations. Ectomycorrhizae have been reported from a number of Australian Gnaphalieae (Warcup 1990), but there is no recent work on thee plants. The oomycete Pustula, a white blister rust, is found quite widely on Asteraceae, with a few occurrences on Goodeniaceae, Araliaceae (Trachymene) and Gentianaceae (Ploch et al. 2010b).

Genes & Genomes. For genome duplications, see Barker et al. (2008) and Schranz et al. (2012). There was an early palaeopolyploidy event involving most or all of the family, and again near the base of Asteroideae and within Mutisioideae. The pattern of duplicate gene retention is distinctive - structural/cell organization genes, but fewer regulatory genes were retained (Barker et al. 2008). A large inversion in the chloroplast genome occurs in most of the family, but not Barnadesioideae and other Asterales (Jansen & Palmer 1987; Bremer 1987; see also below). For satellite DNA diversification in Cardueae, see del Bosque et al. (2014).

Relatively common and wide (with respect to both taxonomy and current geography) hybridisation in Asteroideae in particular is shown by incongruence between topologies based on different genomes, and this makes life for those involved in phylogeny reconstruction rather interesting (e.g. Fehrer et al. 2007; Pelser et al. 2008, 2010, 2012; Soejima et al. 2008; Morgan et al. 2009; Montes-Moreno et al. 2010; Schilling 2011; Smissen et al. 2011; Calvo et al. 2013; Galbany-Casals et al. 2014). Thus there is significant incongruence between relationships suggested by plastid and nuclear sequences in Senecioneae, probably due to ancient hybridisation rather than incomplete lineage sorting (Pelser et al. 2010). Smissen et al. (2011; see also Galbany-Casals et al. 2014) suggest that complex allopolyploidy may have been involved in the origin of at least four clades in Gnaphalieae, one of which is now globally distributed and that together encompass more than half the ca 1,240 species of the tribe. For some hybridization between members of different subtribes in Cichorioideae, see Y. Liu et al. (2013).

Vallès et al (2012) discuss polyploidy and its connection with genome size, etc., while Vallès et al. (2013) summarize genome size variation in the family, although unfortunately little is known about genome size in the basal pectinations. For the reduction of chromosome number in Gnaphalieae, e.g. from n = 12 to n = 3 in Podolepis, see Smissen et al. (2011 and references).

Economic Importance. Timme et al. (2007) provide a phylogeny of the important genus Helianthus; Simpson (2009) summarised what is known of the otherwise rather slight economic importance of the family.

Chemistry, Morphology, etc. For a general entry into the literature of Asteraceae, see papers in Funk et al. (2009a) - there is a helpful glossary; Anderberg et al. (2006) also summarize the variation in the family.

There are tens of thousands of secondary metabolites produced by the family (Calabria et al. 2009 for a convenient summary and entry into the literature), although nothing seems to be known about the secondary chemistry of Hecastocleioideae and Gymnarrhenoideae. See Seaman (1982) for sesquiterpene lactones, Zidara (2008) for those of Cichorieae, Seaman et al. (1990) for diterpenes, Aniszewski (2007) for alkaloids, and Bohm and Stuessy (2001: family) and Sareedenchai and Zidorn (2010: Cichorieae) for flavonoid chemistry; Calabria et al. (2007) produced a phytochemical phylogeny at the tribal level.

The distinctive capitulate inflorescence of Asteraceae lacks a terminal flower and is basically racemose, unlike in its sister taxon, Calyceraceae (Pozner et al. 2012). In Gorteria the outermost (ray) flowers develop both centrifugally and more slowly than the acropetally-developing more central (disc) flowers (Philipson 1953, Harris 1995 and references; Leins & Erbar 2003b; Thomas et al. 2009), and there are several cases where the peripheral flowers develop more or less centrifugally (literature summarized by Pozner et al. 2012). Interestingly, those Calyceraceae which have an inflorescence most similar to that of Asteraceae, and those Asteraceae with some centrifugal development of flowers that can perhaps be linked with the cymose part inflorescences common in the outgroups to Asteraceae, are both derived within their respective families (c.f. Pozner et al. 2012). Similarly, there are floral bracts in some Asteroideae-Heliantheae (the tribe includes taxa that were in Eupatorieae), but they seem to have been reacquired more than once and are certainly not plesiomorphic in the family as was once thought (see e.g. Harris 1995 for literature).

The pappus (modified calyx) is often not the first part of the flower to be initiated, and it may start to develop well after the corolla (see Mukherjee & Harris 1995, Nordenstam 2008), although otherwise the sequence of initiation of flower parts is as might be expected. In general, the very different adult floral morphologies are quite similar early in development (Harris 1995). A corolla ring primordium may initate first, or petals may be initiated separately, i.e., there is variation between early and late corolla tube development, the former perhaps being derived (Harris 1995: Leins & Erbar 2000; Erbar & Leins 2000 for floral development). CYC-like genes appear to be expressed in the abaxial petals here, rather than in the adaxial petals, as in other core eudicots (Citerne et al. 2010).

Floret morphology varies extensively in Asteraceae, and this needs to be put in the context of a tree. Koch (1930 and references) and Manilal (1971) discussed corolla venation; the corolla of the ray flowers of some Asteroideae may even be unvascularized. Many Asteroideae have three-toothed ray florets that give the appearance of being slit-monosymmetric (0:5), but they may be a modified 2:3 bilabiate corolla in which the adaxial lobes have been suppressed (Weberling 1989; Gillies et al. 2002); true slit-monosymmetric flowers are uncommon here. Some taxa, including Barnadesioideae, have a midvein in the petal (see also Carlquist 1976; Gustafsson 1995); corolla variation in Barnesdeioideae alone is extensive (Stuessy & Urtubey 2006). The corolla hairs of Barnadesioideae are distinctive: The epidermal cell is undistinguished, the basal cell is short and thick-walled, and the other cell is longer and has thin walls. There is variation in the orientation of the gynoecium and style branches: carpels superposed, style branches arranged radially to the head surface; carpels collateral, style branches tangential to head surface: see Robinson 1984); details of the distribution of this feature are unknown. Buphthalmum has a hollow style (Leins 2000); I do not know how widespread such styles are in Asteraceae.

There is much variation in pollen morphology in the family, and this has often been described in terms of pollen "types". However, Blackmore et al. (2009) decompose these types into a number of individually-varying characters, and then discuss the distribution of some of these characters across the Asteraceae tree. Caveate pollen is found in Arctoteae and some Lactuceae, and may be more basal on the tree than it is placed here (as a synapomorphy for [Corymboideae + Asteroideae]). Indeed, Blackmore et al. (1984) noted that caveae are evident early in development in Gerbera (Mutisieae), but not later, and suggested that pollen grains of Asteraceae might all be basically caveate - however, the very definition of caveate is unclear (Blackmore et al. 2009). Wortley et al. (2007b) used distinctive pollen characters to help place some genera whose relationships had previously been unclear. There is considerable variation in tapetum "types" in the family (Pacini 1996). See also Skvarla et al. (1977: pollen terminology in Asteraceae), Roque and Silvestre-Capelato (2001: pollen of Gochnatioideae), Wortley et al. (2008: Arctotidae-Cichoridoideae), Wortley et al. (2009: comprehensive bibliography of palynological work in the whole family), Tellería and Katinas (2009: Mutisia), Osman (2009: Cichorioideae-Cardueae), Hong Wang et al. (2009b and references: Cichorioideae-Cichorieae).

The seed coat is poorly developed, as might be expected; there may be calcium oxalate crystals in the inner layer (Guignard 1893). Guignard (1893) also suggested a), that the ovules may be vascularized, and b), that there is sometimes an antiraphe, as in Centaurea. Asteroideae-Heliantheae have distinctive black fruits that are covered by phytomelan (see Graven et al. 1998 for what is known about this compound); they are also described as being carbonized. Nuclear endosperm is sometimes mentioned as being the only endosperm condition found in the family or as a synapomorphy for it (e.g. Tobe & Morin 1996; Inoue & Tobe 1999), but there is in fact considerable variation in endosperm development (Dahlgren 1920; Johri et al. 1992 for references). The embryo of Syneilesis may lack cotyledons entirely (Teppner 2001). For fatty acids in the seeds, see Bamai and Patil (1981). For embryo sac development, which I have not thought about, see e.g. Fagerlind (1939c): there are embryo sacs other than the common monosporic 8-nucleate type.

Phylogeny. There has been much phylogenetic work on Asteraceae, and only a few references are included. Panero and Funk (2002, especially 2008; see also Funk et al. 2005, a supertree, 2009c, a metatree) and Panero et al. (2014) present the phylogeny reflected in the classification above. A large inversion in the chloroplast genome occurs in most of the family, but not Barnadesioideae and other Asterales, and its discovery in the early days of molecular systematics was very exciting, in part because it was consistent with a morphological phylogeny that came out at about the same time (c.f. Jansen & Palmer 1987; Bremer 1987; see also Y.-D. Kim & Jansen 1995; K.-J. Kim et al. 2005; Timme et al. 2005). Much, but not all, of the uncertainty in relationships around the old Mutisioideae seems to have been resolved, with distinctive gene deletions and insertions characterising a number of the clades (Panero & Funk 2008). Morphological data suggested to Roque and Funk (2013: c.f. character states) that Wunderlichioideae and Stifftioideae might form a clade, and there is also some molecular support for this clade (Funk et al. 2014). In the latter study, there was still weaker support for a [Mutisioideae + Stifftiodeae] clade and Perytoideae were outside a [Cichorioidae + Carduoideae] clade, but the latter areas were not the focus of the study and so sampling was poor (Funk et al. 2014). Gymnarrhena was excluded from Asteroideae by Anderberg et al. (2005).

With the realization that the recently-rediscovered Famatinanthus was rather different from other Mutisioideae-Onoserideae, Panero et al. (2014: 14 loci, all chloroplast) examined basal relationships in the family. Famatinanthus was strongly supported as sister to the rest of the family bar Barnadesioideae, and Mutisioideae was the next branch, also with strong support, although support for the Monophyly of Mutisioideae themselves could be better. Support for the positions of the next three clades up was also quite good (Panero et al. 2014). Taking a rather different tack, Mandel et al. (2014) looked at a massive conserved orthologous set of nuclear genes, and found that Centrapalus pauciflorus had sprung outside the other Cichorioideae examined, however, sampling was poor.

For phylogenetic relationships within Barnadesioideae, see Urtubey and Stuessy (2001) and in particular Gustaffson et al. (2001), Gruenstaeudl et al. (2009) and Funk and Roque (2011). The position of Schlechtendalia is uncertain; [Huarpea + Barnadesia may be sister to the rest of the family. For corolla morphology in this subfamily, see Stuessy and Urtubey (2006).

For a phylogeny of Carduoideae-Cardueae, see Garcia-Jacas et al. (2002), Susanna et al. (2006) and Park and Potter (2013). Carduinae are paraphyletic, but for a phylogeny of the monophyletic Centaureinae, see Garcia-Jacas et al. (2001) and Hellwig (2004). For relationships in Echinops, see Garnatje et al. (2005: sectional classification) and Sánchez-Jimenéz et al. (2010), and for those within Cousinia (biphyletic) and relatives, see López-Vinyallonga et al. (2009).

Cichorioideae: Warionia may be sister to all other Cichorieae (Kilian et al. 2009); although no flavonoids have been reported from this tribe, they are diverse in the rest of the tribe (Sareendenchai & Zidorn 2010 - see Zidorn 2008 for sesquiterpene lactones there). Subtribes are monophyletic, although Faberia seems to be a hybrid between a member of Crepidinae and Lactucinae (it looks more like the latter), but relationships between them are only partly resolved (Y. Liu et al. 2013). In Sonchinae, Sonchus is para/polyphyletic (Kim et al. 2007), with woody, island-dwelling forms being independently derived within the clade; for relationships within Scorzonerinae, see Mavroidiev et al. (2004). The classic studies by Babcock (e.g. 1947) on Crepis that assumed that evolution - in this case of the karyotype in particular - was unidirectional need re-evaluation (Enke & Gemeinholzer 2008). Species limits around here are difficult because of apomixis; see Gottschlich (2009) for the complexities of variation in Hieracium in a smallish area of Italy. For a phylogeny of Tragopogon and its relatives, see Mavrodiev et al. (2005) and Bell et al. (2012b), and of the African Gorteriinae, see Funk and Chan (2008). For a phylogeny of Vernonia (Vernonieae), a genus whose circumscription is problematic - either it is huge, or quite small - see Keeley et al. (2007), and for that of the largely Peruvian Liabeae, sister to Vernonieae, see Funk et al. (2012).

Asteroideae; For a phylogeny of Anthemidae in the southern hemisphere, see Himmelreich et al. (2008 and references), Cotula is not monophyletic (Himmelreich et al. 2012) and will probably need to be expanded, and there is extreme polyploidy in Leptinella in particular (Himmelreich et al. 2014). For a delimitation of Anthemis itself, see Lo Presti et al. (2010); for circum-Mediterranean Anthemidae, also their biogeography, see Oberprieler (2005), for Chrysanthemum and other Anthemidae, see Zhao et al. (2010), and for Tanacetum, see Sonboli et al. (2012: little resolution). For relationships within and the evolution of Artemisia, see Vallès et al. (2003) Vallès and Garnatje (2005), Sanz et al. (2008), Pellicer et al. (2010b: genome size, etc., 2011), Garcia et al. (2011: North American taxa) and Riggins and Siegler (2012: paraphyly, etc.). North American Astereae are monophyletic and largely herbaceous (Noyes & Rieseberg 1999), however, Aster is extensively para/polyphyletic, and its limits are now restricted (e.g. Li et al. 2012); for Machaerantherinae, see Morgan et al. (2009). For the Hinterhubera group, see Karaman-Castro and Urbatsch (2009: groupings geographic); Olearia is likely to be polyphyletic (Cross et al. 2002). Strijk et al. (2012) find that Psiadia (and Conyza) are also polyphyletic. Within Gnaphalieae, morphological variation is analyzed by Anderberg (1991a), and phylogenetic relationships there have begun to be disentangled by Bayer et al. (2000). Helichrysum is polyphyletic (Galbany-Casals et al. 2004, 2009, 2010, 2014; Bergh & Linder 2009); Ward et al. (2009), Montes-Moreno et al. (2010), Nie et al. (2013: Anaphalis embedded in Helichrysum) and Bengtson et al. (2014 and references: the South African Metalasia, also diversifcation) further clarify relationships in this tribe; there has been hybridisation (see above). For the phylogeny of the helenioid Heliantheae, see Baldwin et al. (2002), and for that of the iconic Hawaiian silverswords and their island relatives, and of their relatives, the west North America tarweeds (Helenieae-Madiinae), see Baldwin and Wessa (2000) and Carlquist et al. (2004, also Madiinae Showcase). Bidens on Hawaii also shows much variability in growth form, etc., but there is little molecular variation or genetic barriers between the species (Ganders et al. 2000). For the phylogeny of Inuleae (inc. Plucheeae), see Anderberg et al. (2005) and Englund et al. (2009); morphological relationships in Inuleae and Plucheeae are discussed by Anderberg (1991b, c). The sections of Blumea need overhaul, see Pornpongrungrueng et al. (2009). Relationships within Senecioneae, particularly the huge genus Senecio, are beginning to be disentangled (Pelser et al. 2006, esp. 2007, see also Pelser et al. 2010; Calvo et al. 2013); J.-Q. Liu et al. (2006) discuss relationships in the large Ligularia–Cremanthodium–Parasenecio clade, a major part of the Senecioneae-Tussilagininae. For relationships within Euryops, see Devos et al. (2010). For Symphyotrichum and relatives, see Vaezi and Brouillet (2009).

Classification. See Panero and Funk (2008) for the subfamilial classification above (there are, of course, alternative classifications - e.g. Jeffrey 2004); it is similar in basic structure to that in Funk et al. (2009b: as 43 tribes, see the accounts there).

Anderberg et al. (2006) enumerated the genera in the family. However, generic limits in many places are in a state of flux. Vernonia is a classic case of uncertainty - should it include 800-1000 species, or should these species be placed in 20 subtribes, of which two thirds of the genera are mono- or ditypic? (Keeley et al. 2007 for a phylogeny; Robinson 2006 and references for genera). Should there be lumping or splitting in the even larger genus Senecio (see Pelser et al. 2006, esp. 2007)? In the latter case, Senecio species are found in over eight clades, but even so Senecio s. str., at ca 1,000 species, is paraphyletic (Pelser et al. 2007). Substantial adjustments to generic limits will also be needed in Asteroideae-Inuleae-Inulinae (Englund et al. 2009) and in Astereae (e.g. Li et al. 2012). Genera in the Gochnatia area have been split, despite protestations: "we simply seek to insure [sic] that our classifications reflect what we know about evolution" (Funk et al. 2014: p. 879). Finally, hybridisation (see above) makes some genera and even subtribes non-monophyletic. A suggested nomenclatural solution for the area around Helichrysum sounds like a slowly-unfolding disaster, but the alternative, a Helichrysum s.l., seems little better (Galbany-Casals et al. 2014).

Botanical Trivia.Arctium lappa (burdock) infructescences became attached to the dog of a Swiss engineer, George de Mestral, in 1945, and the result was the development of velcro (Wikipedia 2009).